TROPICAL FOREST COMMUNITY ECOLOGY TROPICAL FOREST COMMUNITY ECOLOGY Editors Walter P. Carson University of Pittsburgh Department of Biological Sciences Pittsburgh, PA USA Stefan A. Schnitzer University of Wisconsin – Milwaukee Department of Biological Sciences Milwaukee, WI USA and Smithsonian Tropical Research Institute Apartado 0843-03092, Balboa Republic of Panama This edition first published 2008 ©2008 by Blackwell Publishing Ltd Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell.
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Set in 9/11 pt Photina by Newgen Imaging Systems Pvt. Ltd, Chennai, India Printed in Singapore by Ho Printing Pte Ltd 1 2008 To my parents Walter and Alice Carson: thank you for years of unconditional love and support, and to my son Chris Carson: you have changed the way I see the forest. Carson Contents Preface x Walter P. Carson and Stefan A.
Schnitzer Foreword xi S. Joseph Wright List of Contributors xiii SECTION 1 INTRODUCTION 1 1 Scope of the Book and Key Contributions 3 Stefan A. Schnitzer and Walter P. Carson SECTION 2 LARGE-SCALE PATTERNS IN TROPICAL COMMUNITIES 9 2 Spatial Variation in Tree Species Composition Across Tropical Forests: Pattern and Process 11 Jérôme Chave 3 The Disparity in Tree Species Richness among Tropical, Temperate, and Boreal Biomes: The Geographic Area and Age Hypothesis 31 Paul V.
Ree, and Robyn J. Burnham 4 Explaining Geographic Range Size by Species Age: A Test Using Neotropical Piper Species 46 John R. Paul and Stephen J. Tonsor 5 Patterns of Herbivory and Defense in Tropical Dry and Rain Forests 63 Rodolfo Dirzo and Karina Boege 6 Ecological Organization, Biogeography, and the Phylogenetic Structure of Tropical Forest Tree Communities 79 Campbell O.
Cannon, and Stuart J. Davies 7 Large Tropical Forest Dynamics Plots: Testing Explanations for the Maintenance of Species Diversity 98 Jess K. Zimmerman, Jill Thompson, and Nicholas Brokaw SECTION 3 TESTING THEORIES OF FOREST REGENERATION AND THE MAINTENANCE OF SPECIES DIVERSITY 119 8 Tropical Forest Ecology: Sterile or Virgin for Theoreticians? 121 Egbert G. Leigh, Jr 9 Approaching Ecological Complexity from the Perspective of Symmetric Neutral Theory 143 Stephen P.
Hubbell viii Contents 10 Functional Basis for Resource Niche Partitioning by Tropical Trees 160 Kaoru Kitajima and Lourens Poorter 11 Colonization-related Trade-offs in Tropical Forests and Their Role in the Maintenance of Plant Species Diversity 182 Helene C. Muller-Landau 12 Treefall Gaps and the Maintenance of Plant Species Diversity in Tropical Forests 196 Stefan A. Schnitzer, Joseph Mascaro, and Walter P. Carson 13 Challenges Associated with Testing and Falsifying the Janzen–Connell Hypothesis: A Review and Critique 210 Walter P.
Leigh, Jr, and Stefan A. Schnitzer 14 Seed Limitation and the Coexistence of Pioneer Tree Species 242 James W. Dalling and Robert John 15 Endophytic Fungi: Hidden Components of Tropical Community Ecology 254 A. Elizabeth Arnold SECTION 4 ANIMAL COMMUNITY ECOLOGY AND TROPHIC INTERACTIONS 273 16 Tropical Tritrophic Interactions: Nasty Hosts and Ubiquitous Cascades 275 Lee A.
Dyer 17 Variation in Tree Seedling and Arbuscular Mycorrhizal Fungal Spore Responses to the Exclusion of Terrestrial Vertebrates: Implications for How Vertebrates Structure Tropical Communities 294 Tad C. Theimer and Catherine A. Gehring 18 Ecosystem Decay in Closed Forest Fragments 308 John Terborgh and Kenneth Feeley 19 Resource Limitation of Insular Animals: Causes and Consequences 322 Gregory H. Adler 20 Tropical Arboreal Ants: Linking Nutrition to Roles in Rainforest Ecosystems 334 Diane W.
Davidson and Steven C. Cook 21 Soil Fertility and Arboreal Mammal Biomass in Tropical Forests 349 Carlos A. Peres SECTION 5 SECONDARY FOREST SUCCESSION, DYNAMICS, AND INVASION 365 22 Processes Constraining Woody Species Succession on Abandoned Pastures in the Tropics: On the Relevance of Temperate Models of Succession 367 Chris J. Peterson and Walter P.
Carson 23 Chance and Determinism in Tropical Forest Succession 384 Robin L. Chazdon 24 Exotic Plant Invasions in Tropical Forests: Patterns and Hypotheses 409 Julie S. Denslow and Saara J. DeWalt Contents ix SECTION 6 TROPICAL FOREST CONSERVATION 427 25 Linking Insights from Ecological Research with Bioprospecting to Promote Conservation, Enhance Research Capacity, and Provide Economic Uses of Biodiversity 429 Thomas A.
Capson, Luis Cubilla-Rios, Daniel A. Emmen, William Gerwick, Mahabir P. Heller, Kerry McPhail, Eduardo Ortega-Barría, Dora I. Solis, and Phyllis D.
Coley 26 Tropical Rainforest Conservation: A Global Perspective 442 Richard T. Corlett and Richard B. Primack 27 Environmental Promise and Peril in the Amazon 458 William F. Laurance 28 Contributions of Ecologists to Tropical Forest Conservation 474 Francis E.
Putz and Pieter A. Zuidema Index 491 Preface It is not hyperbole to say that there has been ecological organization. Indeed, it is difficult to an explosion of research on tropical forest ecol- understand communities without understanding ogy over the past few decades. The establishment the ecology of populations and individuals.
We of large forest dynamics plots in tropical forests decided to focus on forest communities because, worldwide, in and of itself, has led to a near rev- to date, that is where the bulk of research on trop- olution in our understanding of forest change. In ical community ecology has been conducted. We addition, there has been a substantial increase in acknowledge that our focus has forced us to omit the use of models and experiments to test long- many important studies. Nonetheless, the empha- standing theories developed to explain the striking sis on tropical forest community ecology provides patterns found in tropical forests and the putative enough material to fill multiple edited volumes, mechanisms that underlie these patterns.
When and thus we have attempted to focus on the areas we started this project, we felt that a compre- that have received the most empirical attention, hensive synthesis of tropical forest community along with some topics that are currently nascent, ecology was necessary in order to help the field but are rapidly becoming key areas in tropical move forward. Of course, no single volume could ecology. Nonetheless, this book is our attempt to Each chapter in this book was reviewed by at make a significant contribution to the field, and to least two relevant experts. We thank these review- ask anew: What are the main theories in tropical ers for their efforts and we are indebted to all of ecology, and which ones are supported or refuted them.
We will not list them by name, thus allow- by empirical data? Thus, we have attempted to ing them to remain anonymous. We also thank the assemble a volume that describes the most up-to- production team at Newgen Imaging Systems, and date findings on the important theories of tropical our editors at Blackwell for guiding us through the forest community ecology. We hope that this book publication process. accomplishes this goal to the degree possible, while This book, as with all edited volumes, would at the same time providing a road map of what we not have been possible without the dedicated con- know, what we think we know, and where future tributions of the authors, each of whom is an research is most needed.
expert in his or her respective area of study. For The focus of the chapters in the volume is at their hard work, truly top-notch contributions, the community level because this is where some and their patience throughout this process, we of the most fundamental questions in tropical owe them a great deal of gratitude. This book is a ecology exist. Indeed, perhaps the greatest chal- tribute to their research, along with the research lenge to community ecologist is to explain what of all of the other scientists whose work is cited in processes account for the maintenance of the stag- this volume.
gering diversity of plants and animals common in Walter P. Carson tropical forests around the globe. Still, our empha- Pittsburgh, Pennsylvania sis on communities definitely reflects our bias as 2007 community ecologists. While we have focused on communities, we certainly recognize the impor- Stefan A.
Schnitzer tant contributions to tropical ecology that have Gamboa, Republic of Panama come from those who study different levels of 2007 Foreword The present volume captures the excitement gen- forest communities are structured and are only erated by an explosion in tropical forest research. now beginning to be explored. When I was a graduate student in the late 1970s, In the 1970s, we would have been mysti- it seemed to be possible to read every new arti- fied by functional (Chapter 10) and phylogenetic cle published on tropical forests. The ISI Web of (Chapter 20) approaches to plant community Science© confirms this schoolboy memory.
Just ecology and the knowledge base in physiology, 289 articles published between 1975 and 1979 morphology and molecular genetics that makes included the words “forest*” (for forest, forested these approaches possible today. Both approaches or forests) and the name of a tropical country have the potential to reduce the immense number (or tropic*) in their titles. By reading just one or of species of tropical forest plants to a man- two articles a week, I was able to keep abreast ageable number of ecologically distinct groups of the entire literature on tropical forests. This or crucial relationships among species’ traits.
would be nearly impossible today. Between 2002 Today, we are striving to bring functional, phy- and 2006, 2593 articles met the criteria described logenetic and ecological approaches together for above reflecting a nine-fold increase in the rate 6000 plus tropical tree species found in the of publication of tropical forest articles since the network of large Forest Dynamics Plots main- late 1970s. This explosion has been driven by tained by the Center for Tropical Forest Science new discovery; new theory; new technology; new (Chapter 7). challenges posed by global change, deforestation A graduate student in the late 1970s would and other threats to tropical biodiversity; and have been familiar with the plant favorable- ongoing interest in theory posed in the 1970s ness (Chapters 3 and 4), regeneration niche and earlier.
This volume illustrates each of these (Chapter 6), Janzen–Connell (Chapter 13) and developments.